903 resultados para Plasticity
Resumo:
We describe a first-principles-based strategy to predict the macroscopic toughness of a gamma-Ni(Al)/alpha-Al2O3 interface. Density functional theory calculations are used to ascertain energy changes upon displacing the two materials adjacent to the interface, with relaxation conducted over all atoms located within adjoining rows. Traction/displacernent curves are obtained from derivatives of the energy. Calculations are performed in mode I (opening), mode II (shear) and at a phase angle of 45 degrees. The shear calculations are conducted for displacements along < 110 > and < 112 > of the Ni lattice. A generalized interface potential function is used to characterize the results. Initial fitting to both the shear and normal stress results is required to calibrate the unknowns. Thereafter, consistency is established by using the potential to predict other traction quantities. The potential is incorporated as a traction/displacement function within a cohesive zone model and used to predict the steady-state toughness of the interface. For this purpose, the plasticity of the Ni alloy must be known, including the plasticity length scale. Measurements obtained for a gamma-Ni superalloy are used and the toughness predicted over the full range of mode mixity. Additional results for a range of alloys are used to demonstrate the influences of yield strength and length scale.
Resumo:
青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障,经过近六十年的采伐后,取而代之的是大量人工种植的云杉纯林。目前,这些人工林已经表现出树种单一,结构层次简单等生态问题,其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性,完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分,对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此,研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象,选择林下11种具有不同喜光特性的常见植物,分别设置人工林林冠下及成熟林窗为研究样地,通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析,研究林下植物对不同光生境的适应策略及其适应能力,揭示不同物种对人工林生境的适应共性,为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下: 在叶片形态和物质分配特征方面:在林窗光生境中,11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时,林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加,这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后,植物可能更多地采取调节叶片组织细胞水平(即叶片功能细胞形态)及叶片器官水平(即单个叶片形态)特征的策略来适应各类生境,而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面:多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多,而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面:在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下,多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率,并在中午12:00~14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境,即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境,通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之,植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关,但另一些参数(Fv/Fm日变化、色素含量及比例、叶氮相对含量)受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面:在叶片各表型参数中,器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性;叶片光合组织的可塑性大于非光合组织可塑性;反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合,即喜光种及轻度耐荫种各参数可塑性最高,深度耐荫种可塑性最小,而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面:喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低,深度耐荫种疏花槭却恰恰相反,更多数物种(7种植物)在两种生境中密度变化趋势不明显。从分布格局来看,7种植物在两种生境中均为聚集分布,但聚集强度为林窗>林内;少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型,在林内生境中的分布型发生改变而成为随机型,说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关,同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.
Resumo:
在青藏高原东部的亚高山针叶林区,如何尽快恢复这一生态脆弱地区的植被,改变生态环境恶化的趋势,是一个十分重要的课题。光一直被认为是植物种间相互替代,尤其是森林演替过程中植物相互替代或植被恢复中的关键环境要素之一。植物能否适应林冠下或林窗中异质的、或多变的光照条件,对其在林中的生存、分布、更新以及森林动态都是非常重要的。 本文以青藏高原东部亚高山针叶林的主要森林类型——岷江冷杉林群落的几种树苗为研究对象,采用实验生态学、生理及生物化学等方法,通过模拟针叶林不同大小林窗内光照强度的变化,在中国科学院茂县生态站内采用遮荫处理设置6个光照梯度(100、55、40、25、15与7%全光照),来研究具有不同喜光特性的植物对光强的响应与适应机制,其研究结果可为揭示亚高山针叶林的演替规律、以及人工林下幼苗的存活与定居提供科学依据,也能为苗木的生产与管理提供科学指导,尤其是对针阔树种在不同光强下的响应与适应的比较研究,能为如何将阔叶树种整合到人工针叶林中提供新的思路。 光强对植物生长的影响 光强对植物的生长具有重要作用,不同植物在各自适宜的光强梯度下才能生长良好。通过一个野外盆栽实验,来研究不同光强对植物生长的影响(第三章)。主要研究结果如下,低光强下植物株高/茎生物量增加,说明植物会将生物量更多用于高生长,以便有效地拦截光资源;在强光下,植物将生物量更多地向根部分配,使得植物在强光下能够吸收更多的水分,而避免干旱胁迫。 在第一个生长季节,以相对生长速率(RGR)表示,红桦和青榨槭在100%全光照下RGR最大,粗枝云杉在55%最大,岷江冷杉在25-40%下较好;然而,在第二个生长季节,2种阔叶树的相对生长速率(RGR)的适宜光强则变为25-55%,云杉为55-100%,而冷杉为25-100%。可见,从第一年到第二年,2种阔叶树苗更适宜在部分荫蔽的条件下生长;而2种针叶树苗对光的需求则逐渐增加,这可能是增加对根生物量相对投资的结果,因为以这种方式,强光下生长的针叶树幼苗更能保持其内部水分平衡,其生长不会因干旱胁迫而受到严重影响。另外,严重遮荫会引起冷杉幼苗死亡。 植物对光强的生理适应 植物可以通过自身形态和生理特征的调整,来发展不同的光能利用策略从而能够在林中共存。通过一个野外盆栽实验,研究了不同光强下生长的几种树苗的生理特征(第四章)对不同光强的响应与适应。结果显示:强光下,粗枝云杉和红桦的光合能力增加,而岷江冷杉和青榨槭在中度遮荫(25-55%)的条件下光合能力最大。植物叶氮和叶绿素含量增高,而光补偿点和暗呼吸速率降低,这些都是植物对低光环境的适应性反应;而强光下植物叶片和栅栏组织变厚,是对强光的一种保护性反应。 植物对光的可塑性反应 不同植物会表现出对光适应有利的生理和形态可塑性反应。本文对第三章、第四章的实验数据进行可塑性指数分析,来研究植物对光强的表型可塑性反应(第五章)。结果显示,生理特征调整是植物对不同光环境的主要适应途径。红桦和青榨槭的可塑性指数平均值要大于粗枝云杉和岷江冷杉,充分表明这2种阔叶树在生理和形态上较强的可塑性更有利于对光环境的适应,而具有比耐荫树种更强的适应能力。另外,2种针叶树相比,云杉的适应性更强。本研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说。 植物的光抑制与防御 当植物叶片吸收了过多光能,会发生光抑制现象。植物对光抑制的敏感性及防御能力对其生长具有重要意义。本文通过两个野外盆栽实验,研究了生长在强光下(第六章)和变化光强下(第八章)植物的光抑制现象及其防御策略。结果表明,在强光下或从遮荫状态转入强光下,植物都会发生光抑制,其对光抑制的敏感性与植物的耐荫性(或喜光)和演替状态有密切联系。长期生长在强光下的植物受到光抑制是可恢复的,而当处于荫蔽环境的植物突然暴露于强光下时,受到的光抑制不能完全恢复,可能是(部分)光合机构受到破坏的缘故。粗枝云杉和青榨槭防御光抑制伤害的能力较强,热耗散是其防御光抑制的主要途径。长期的强光作用能使岷江冷杉和红桦发生严重光抑制,甚至光伤害,而红桦能够通过“凋落老叶,萌发新叶”的途径来适应新的强光环境。 How to restore the vegetation of subalpine coniferous forest in eastern Qinghai-Tibet Plateau, and change the trend of ecological deterioration is a very important issue. Acclimation of tree seedlings to different and varing light environment affects to a great extent the successful regeneration and establishment of subalpine coniferous forests in southwestern China’s montane forest areas, because the ability to respond to such changing resource are commonly assumed to be critical to plant success, and have a growth advantage than others. In this paper, several species seedlings in Abies faxoniana community were chosed to study the response and adaptation to light intensity and the interspecific differences of adaptability in six shaded sheds (100, 55, 40, 25, 15 and 7% of full sunlight) in the Maoxian Ecological Station of Chinese Academy of Sciences. Our results could provide a strong theoretical evidence for understanding the forest succession laws of subalpine coniferous forests, and the survival and settlement of seedlings under plantations, and provide scientific direction for the production and management of seedlings, especially the comparative studies of the acclimation to light between the conifer and broadleaf trees could provide new ideas for how to integrate the broad-leaved trees into the artificial coniferous forest. Growth under different light intensity Light intensity plays an important role on plant growth. One field experiments was conducted to study the growth of tree seedlings of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii under different light intensities. The results showed that plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources, while biomass greater allocation to the roots, could make plants under high light environment absorb more water, and avoid drought stress. During the first growing season, the relative growth rates (RGRs) of Betula albo-sinensis and Acer davidii had the greatest values under the 100% of full light, for 55% of Picea asperata, and for 25-40% of Abies faxoniana. However, in the second growing season the the relative growth rates of the two broad-leaved trees changed and were appropriate for 25-55% of full light, for 55-100% of spruce, and for 25-100% of fir. Thus, from the first year to the second year, two broad-leaved seedlings maybe more suitable to partly shading environment, and two coniferous seedlings would have an increase in light demand, which may be an increased root biomass investment. Because in this way, seedlings grown under high light could better maintain their internal water balance, and thus its growth would not be seriously affected by drought stress. In addition, serious shading would cause fir seedlings to die. Acclimation of physiology to light Plants could coexist in forest ecosystem by forming different strategies of light use. One field experiments was conducted to study the acclimation of tree seedlings to different light intensity of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii. The results showed that the photosynthetic capacity of Picea asperata and Betula albo-sinensis exhibited a general tendency of increase with more light availability; but for Abies faxoniana and Acer davidii seedlings, their highest values of the same parameters were found under intermediate light regime (i.e. 25-55% of PFD relative to full sunlight). Plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources. Leaf nitrogen and chlorophyll content increased, while dark respiration rate and light compensation points decreased, all of which were adaptive response to the low light environment. On the contrary, plants under high light environment had the thicken leaves and palisade tissue, which was a protective response to high light. Phenotypic plasticity to light Phenotypic plasticity can be exhibited in morphological and physiological processes. Physiological characteristical adjustment is the main for plant adaptation to different light environment.The means of plasticity indexes for Betula albo-sinensis and Acer davidii seelings were greater than Picea asperata and Abies faxoniana, amplied that the two broad-leaved trees were much more adaptable to the environment. In addition, spruce had the higher adaptablity than fir. The findings supported the hypothesis that the ecological characteristics of the species determined the biological status and its biological habitat selection. Photoinhibition and photoprotection to light Compared with conifer, broad-leaved trees could better change leaf morphology and adjust biomass allocation to adapt to changing light environment. However, excess light can photoinhibit photosynthesis and may lead to photooxidative destruction of the photosynthetic appatus. Two field experiments were conducted to study the photoinhibition of photosynthesis. The results showed that when plants grown under high light environment or plants transferred from low to high irradiance, the four tree seedlings would undergo a period of photoinhibition. In four species, photoinhibited leaves could recover to initial photosynthetic rates when they were long-term planted under high light environment. However, when plants were suddenly exposed to high irradiance, this photoinhibition could not be reversible, may be the photosynthesis apparatus were (or partly) photooxidatively destructed.
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本研究通过粗枝云杉不同种群进行的温室半控制试验,采用植物生态学、生理学和生物化学的研究方法,系统地研究了粗枝云杉不同种群抗旱性的生长、形态、生理和生化机理,并结合有关研究进行综合分析,得出主要研究结论如下: 1.粗枝云杉对干旱胁迫的综合反应 粗枝云杉在干旱胁迫下的适应机制为:(1)相对生长速率及植株结构的调整:干旱胁迫下虽然植株相对生长速率显著降低,且有相对较多的生物量向根部分配,但并未发现细根/总根比增加。(2)粗枝云杉对干旱胁迫的光合作用表现为:干旱胁迫显著地降低了控制的理想条件下的气体交换,但干旱胁迫对PSII最大光化学效率(Fv/Fm)没有影响,表明干旱并未影响到光合机构。(3)干旱还影响了很多生理生化过程,包括渗透调解物质(游离脯氨酸)、膜脂过氧化产物、脱落酸(ABA)含量的增加,以及保护酶活性的升高。这些结果证明植物遭受干旱胁迫后发生了一系列的形态、生理和生化响应,这些变化能提高干旱时期植物的存活和生长能力。 2.粗枝云杉不同种群对干旱胁迫反应的种群差异 粗枝云杉三个种群-干旱种群(四川丹巴和甘肃迭部)和湿润种群(四川黑水)对干旱适应不同,这种不同应归因于它们采用的用水策略不同:在水分良好和干旱胁迫条件下,受试种群在相对生长速率和水分利用效率(WUE)方面都表现出显著的种群间差异。与湿润种群相比,干旱种群在两种水分条件下有更高的WUE。粗枝云杉不同种群的碳同位素组分(δ13C)只在干旱胁迫下有显著差异,并且这种差异在水分良好时比干旱胁迫条件下小,说明生理响应和干旱适应性之间的关系受植物内部抗旱机制和外部环境条件(如水分可利用性)或两者互作效应的影响。这些结果说明干旱种群和湿润种群所采用的用水策略不同。干旱种群有更强的抗旱能力,采用的是节水型的用水策略,而湿润种群抗旱能力较弱,采用的是耗水型的用水策略。 3. 遮荫对粗枝云杉不同种群抗旱性影响 干旱胁迫显著降低了全光条件下叶相对含水量(RWC)、相对生长速率、气体交换参数、PSII的有效量子产量(Y),提高了非光化学猝灭效率(qN)、水分利用效率、脯氨酸(PRO)积累、脱落酸(ABA)含量及保护酶活性。然而这种变化在遮荫条件下不明显。我们得出结论适度遮荫降低了干旱对植物的胁迫作用。另一方面,在干旱条件下,与湿润种群相比,干旱种群抗旱性更强,表现在干旱种群净光合速率与单位重量上叶氮含量(Nmass)降低较少。另外,干旱种群表现出更为敏感的气孔导度,更高的热耗散能力(qN)能力、用水效率、ABA积累、保护酶活性,以及更低的总用水量、相对生长速率。这一结果表明这两种群采用不同的生理策略对干旱和遮荫做出反应。许多生长和生理反应差异与这两个种群原产地气候条件相适应。 4. 外源脱落酸(ABA)喷施对粗枝云杉不同种群抗旱性影响 外源ABA喷施在干旱和水分良好条件下均不同程度地提高了根/茎比,表明根和茎对ABA敏感程度不同。实验结果还表明,外源ABA喷施对这两个种群在干旱胁迫期间影响不同。干旱胁迫期间,伴随着ABA喷施,湿润种群净光合速率(A)显著降低,而干旱种群净光合速率变化不明显。另一方面,外源ABA喷施显著提高了干旱条件下干旱种群的单位叶面积重(LMA)、根/茎比、细根/总根(Ft)比、水分利用效率(WUE)、ABA含量, 以及保护酶活性。然而,外源ABA喷施对湿润种群的上述测定指标没有显著影响。这一结果表明干旱种群对外源ABA喷施更为敏感, 反应在更大的气孔导度降低,更高的生物量可塑性,及更高的水分利用效率、ABA含量和保护酶活性。综上所述,我们得出结论,粗枝云杉对外源ABA敏感性因种群的不同而不同。该研究结果可为两个明显不同种群在适应分化方面提供强有力的证据。 Arid or semi-arid land covers more than half of China's land territory. In arid systems, severe shortages of soil water often coincide with periods of high temperatures and high solar radiation, producing multiple stresses on plant performance. Protection from high radiation loads in shaded microenvironments during drought may compensate for a loss of productivity due to reduced irradiance when water is available. Additionally, ABA, a well-known stress-inducible plant hormone, has long been studied as a potential mediator for induction of drought tolerance in plants. Picea asperata Mast., which is one of the most important tree species used for the production of pulp wood and timber, is a prime reforestation species in western China. In this experiment, different population of P. asperata were used as experiment material to study the adaptability to drought stress and population differences in adaptabiliy, and the effects of shade and exogenous abscisic acid (ABA) application on the drought tolerance. Our results cold provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem in the arid and semi-arid area, and provide a strong evidence for adaptive differentiation of different populations, and so may be used as criteria for species selection and tree improvement. The results are as follows: 1. A large set of parallel response to drought stress Drought stress caused pronounced inhibition of the growth and increased relatively dry matter allocation into the root; drought stress also caused pronounced inhibition of photosynthesis, while drought showed no effects on the maximal quantum yield of PSII photochemistry (Fv/Fm) in dark-adapted leaves, indicating that drought had no effects on the primary photochemistry of PSII. However, in light-adapted leaves, drought reduced the quantum yield of PSII electron transport (Y) and increased the non-photochemical quenching (qN). Drought also affected many physiological and biochemical processes, including increases in superoxide dismutase (SOD), ascorbate peroxidase (APX) activities, malondialdehyde and ABA content. These results demonstrate that there are a large set of parallel changes in the morphological, physiological and biochemical responses when plants are exposed to drought stress; these changes may enhance the capability of plants to survive and grow during drought periods. 2. Difference in adaptation to drought stress between contrasting populations of Picea asperata There were significant population differences in growth, dry matter allocation and water use efficiency. Compared with the wet climate population (Heishui), the dry climate population (Dan ba and Jiebu) showed higher LMA, fine root/total root ratio and water use efficiency under drought-stressed treatments. The results suggested that there were different water-use strategies between the dry population and the wet population. The dry climate population with higher drought tolerance may employ a conservative water-use strategy, whereas the wet climate population with lower drought tolerance may employ a prodigal water-use strategy. These variations in drought responses may be used as criteria for species selection and tree improvement. 3. The effects of shade on the drought tolerance For both populations tested, drought resulted in lower needle relative water content (RWC), relative growth rate (RGR), gas exchange parameters and effective PSII quantum yield (Y), and higher non-photochemical quenching (qN), water use efficiency (WUE), proline (PRO) and abscisic acid (ABA) accumulation, superoxide dismutase (SOD), ascorbate peroxidase (APX) activities as well as malondialdehyde (MDA) levels and electrolyte leakage in sun plants, whereas these changes were not significant in shade plants. Our study results implied that shade, applied together with drought, ameliorated the detrimental effects of drought. On the other hand, compared with the wet climate population, the dry climate population was more tolerant to drought in the sun treatment, as indicated by less decreases in A and mass-based leaf nitrogen content (Nmass), more responsive stomata, greater capacity for non-radiative dissipation of excitation energy as heat (analysed by qN), and higher WUE,higher level of antioxidant enzyme activities,higher ABA accumulation as well as lower MDA content and electrolyte leakage. Many of the differences in growth and physiological responses reported here are consistent with the climatic differences between the locations of the populations of P. asperata. 4. The effects of exogenous abscisic acid (ABA) application on the drought tolerance For both populations tested, exogenous ABA application increased root/shoot ratio (Rs) under well-watered and drought-stressed conditions, indicating that there was differential sensitivity to ABA in the roots and shoots. However, it appeared that ABA application affected the two P. asperata populations very differently during drought. CO2 assimilation rate (A) was significantly decreased in the wet climate population, but only to a minor extent in the dry climate population following ABA application during soil drying. On the other hand, ABA application significantly decreased stomatal conductance (gs), transpiration rate (E) and malondialdehyde (MDA) content, and significantly increased leaf mass per area (LMA), Rs, fine root/total root ratio (Ft), water use efficiency (WUE), ABA contents, superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) activities under drought condition in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet population plants. The results clearly demonstrated that the dry climate population was more responsive to ABA application than the wet climate population, as indicated by the strong stomata closure and by greater plasticity of LMA and biomass allocation, as well as by higher WUE, ABA content and anti-oxidative capacity to defense against oxidative stress, possibly predominantly by APX. We concluded that sensitivity to exogenous ABA application is population dependent in P. asperata. Our results provide strong evidence for adaptive differentiation between populations of P. asperata.
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土壤是人类赖以生存的自然环境和农业生产的重要资源,世界面临的粮食、资源和环境问题与土壤密切相关,目前危害土壤的主要因素是干旱和重金属污染。杨树具有适应性强、生长快和丰产等特性,本论文以青杨组杨树为模式植物,采用植物生态、生理及生物化学等方法,研究杨树对土壤干旱和锰胁迫的生态生理反应以及种群间差异,研究成果可为我国干旱半干旱地区营造人工林、防止沙漠化提供理论依据,也为恢复与重建重金属污染地区退化生态系统提供科学指导。主要研究结果如下: 1. 青海杨不同种群对干旱胁迫的响应差异 干旱胁迫显著降低了两个青海杨种群(干旱种群和湿润种群)生物量积累,包括株高、基径、干物质积累等,通过植物结构的调整,有更多的生物量向根部分配。干旱胁迫还显著降低了叶绿素和类胡萝卜素含量,增加了游离脯氨酸和总氨基酸含量。另一方面,干旱胁迫诱导了活性氧的积累,作为第二信使,激活了抗氧化系统,包括抗坏血酸(ASA)含量和酶系统如超氧化物歧化酶(SOD),愈创木酚过氧化物酶(GPX),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)。这样,杨树既有避旱机制又有耐旱机制,使其在干旱胁迫下有相当程度的可塑性。与湿润种群相比,干旱种群杨树有更多的生物量分配到根部,积累了更多的游离脯氨酸和总氨基酸来进行渗透调节,并且有更有效的抗氧化系统,包括更高含量的ASA 和更高活性的APX 和GR,这些使得干旱种群杨树比湿润种群杨树对干旱有更好的耐性。 2. 喷施硝普钠(SNP)对青海杨阿坝种群干旱胁迫耐性的影响 干旱胁迫显著的降低了青海杨阿坝种群的生长和生物量积累以及叶片相对含水量,还诱导了脯氨酸的合成以进行渗透调节。干旱胁迫下过氧化氢(H2O2)显著累积从而造成对膜脂和蛋白的伤害,使得丙二醛和蛋白羰基含量升高。干旱胁迫下喷施SNP可以减轻干旱胁迫造成的伤害,包括增加叶片的相对含水量,增加脯氨酸和总氨基酸的积累,并激活抗氧化酶系统如SOD,GPX和APX,从而减少丙二醛(MDA)和蛋白羰基(C=O)的积累,但是在水分良好情况下SNP的效果不显著。 3. 青杨不同种群对锰胁迫的生长与形态响应差异 在同一锰浓度下,干旱种群的耐性指数都要高于湿润种群,这表明青杨对干旱和高锰胁迫具有交叉耐性。两个种群的株高,生物量和叶绿素含量都随锰浓度的升高而逐渐下降。就累积浓度而言,0 和0.1 mM 锰胁迫下,干旱种群积累的锰浓度要高于湿润种群,而在高浓度锰胁迫下(0.5 和1 mM),湿润种群要高于干旱种群。在0,0.1 和0.5 mM下,锰大多积累在根中,叶片次之,茎中最少。而在1 mM,锰更多的积累在叶片中。就累积总量而言,在各个锰浓度胁迫下,根,茎和叶相比,两个种群青杨都是叶片累积的锰总量要高于根和茎。两个种群间比较,对照中没有显著区别,0.1 mM 锰胁迫下,湿润种群根中累积的锰要高于干旱种群,而在地上部中,干旱种群要高于湿润种群。而0.5 和1 mM 锰胁迫下,根、叶、茎+叶、根+茎+叶中,锰累积总量都是湿润种群高于干旱种群。锰胁迫下,青杨叶片数和叶面积包括总叶面积和平均叶面积都显著降低。叶片横切面的光学显微观察结果表明未经锰胁迫的栅栏组织的细胞饱满,海绵组织发达、清晰;胁迫后杨树叶片栅栏组织细胞出现不同程度的皱缩,海绵组织几乎不可见,此外还发现输导组织在胁迫下密度变小和分生组织严重割裂等现象。 4. 青杨不同种群对锰胁迫的生理与生化响应差异 青杨两个种群脱落酸(ABA)含量在锰胁迫下都显著增加,干旱种群的增幅更大。三种多胺含量在锰胁迫下显示了不同的响应趋势:腐胺在两个种群的各个锰处理下都增加,亚精胺只在干旱种群中显著增加,而精胺除了干旱种群在1 mM 下有所增加外,在锰胁迫下变化很小。谷胱甘肽含量随锰浓度升高而增加,在0.5 mM 锰时达到最高值,1mM 时有所下降。植物络合素(PCs)含量与非蛋白巯基(NP-SH)趋势相似,随锰浓度的升高而增加,且干旱种群中含量要高于湿润种群。锰处理还引起氧化胁迫,表现为过氧化氢和丙二醛含量增加。SOD 活性在湿润种群中,在0 到0.5 mM 锰胁迫下活性升高,但在1 mM 锰胁迫时,其活性有所下降。而在干旱种群中,SOD 活性变化较小,并始终维持在一个较高的水平。APX 活性在两个种群中都随锰浓度的升高而增加,干旱种群活性要高于湿润种群。锰胁迫还显著增加了酚类物质的含量,同时GPX 和多酚氧化酶(PPO)活性也随锰浓度的升高而增加。干旱种群的酚类含量和GPX 与PPO 活性都要高于湿润种群。锰胁迫还改变了氨基酸的含量和构成,根据锰胁迫下浓度变化的不同,可以将游离氨基酸分为三组:第一组包括,谷氨酸,丙氨酸和天门冬氨酸,这一组氨基酸含量在锰胁迫下有所下降。第二组包括缬氨酸,亮氨酸和苏氨酸含量在锰胁迫下基本不变化或变化很小。剩下的氨基酸为第三组,这组氨基酸含量在锰胁迫下显著增加,而根据增加的幅度又可以将它们分为两个亚组,丝氨酸,酪氨酸,苯丙氨酸,组氨酸和脯氨酸,在1 mM 下的含量是对照的4 倍以上。异亮氨酸,赖氨酸,精氨酸和甘氨酸含量在1 mM 下是对照含量的2 倍以下。同时,同一锰浓度下,干旱种群比湿润种群积累的氨基酸含量要高。 Soil is the indispensable environment for human survival and important resource foragriculture development. Food and environmental problems facing the world are all closelyrelated to soil and nowadays it is threatened by many factors, among which drought stress andheavy metal pollution are the most serious ones. Poplars (Populus spp.) are importantcomponents of ecosystem and suitable as a source of fuel, fiber and lumber due to their fastgrowth. In this study, different populations of Section Tacamahaca spach were used as modelplants to investigate the adaptability to drought stress and manganese toxicity and differencesbetween populations from dry and wet climate regions. Our results can provide theoreticalevidence for the afforestation and prevention of desertification in the arid and semi-arid areas,and also can supply scientific direction for the reconstruction and rehalibitation of ecosystemscontaminated by heavy metals. The results are as follows: 1. Differences in ecophysiological responses to drought stress in two contrastingpopulations of Populus przewalskii Drought stress not only significantly affected dry mass accumulation and allocation, butalso significantly decreased chlorophyll pigment contents and accumulated free proline andtotal amino acids. On the other hand, drought also significantly increased the levels ofabscisic acid and reactive oxygen species, as secondary messengers, to induce the entire set ofantioxidative systems including the increase of reduced ascorbic acid content and the activities of superoxide dismutase, guaiacol peroxidase, ascorbate peroxidase and glutathioneredutase. Thus the combination of drought avoidance and tolerance mechanisms conferredpoplar a high degree of plasticity in response to drought stress. Compared with the wetclimate population, the dry climate population showed lower dry matter accumulation andallocated more biomass to root systems, and accumulated more free proline and total aminoacids for osmotic adjustment. The dry climate population also showed more efficientantioxidant systems with higher content of ascorbic acid and higher activities of ascorbateperoxidase and glutathione redutase than the wet climate population. All these made the dryclimate population superior in adaptation to drought stress than the wet climate population. 2. Effect of exogenous applied SNP on drought tolerance in Populus przewalskii Drought stress significantly increased hydrogen peroxide content and caused oxidativestress to lipids and proteins assessed by the increase in malondialdehyde and total carbonylcontents, respectively. The cuttings of P. przewalskii accumulated proline and other aminoacids for osmotic adjustment to lower water potential, and activated the antioxidant enzymes such as superoxide dismutase, guaiacol peroxidase and ascorbate peroxidase to maintain thebalance of generation and quenching of reactive oxygen species. Moreover, exogenous SNPapplication significantly heightened the growth performance of P. przewalskii cuttings underdrought treatment by promotion of proline accumulation and activation of antioxidant enzymeactivities, while under well-watered treatment the effect of SNP application was very little. 3. Morphological responses to manganese toxicity in the two contrasting populations ofPopulus cathayana High concentration of manganese caused significant decrease in shoot height andbiomass accumulation. The tolerance index of the dry climate population was significantlyhigher than that of the wet climate population, suggesting the superior Mn tolerance in theformer and the existence of cross-tolerance of drought stress and high Mn toxicity. Injuries tothe leaf anatomical features were also found as the reduced thickness in palisade and spongyparenchyma, the decreased density in the conducting tissue and the collapse and split in themeristematic tissue in the central vein. As for the Mn concentrations in the plant tissues, under0, 0.1 and 0.5 mM, most of the Mn accumulated in the roots, then leaves, and stem the least, while under 1 mM, most of the Mn accumulated in the leaves. As far as the total amounts ofMn extraction are concerned, the leaf extracted more Mn than the root and stem in the twopopulations under various Mn concentrations. There is no difference between the twopopulations under control. Under 0.1 mM, the wet climate population extracted higher Mn inthe root than the dry climate population, while in the shoot, the dry climate populationextracted much more Mn. Under 0.5 and 1 mM, the wet climate population translocated moreMn both in the root and the shoot than the dry climate population. 4. Physiological and biochemical responses to manganese toxicity in the two contrastingpopulations of Populus cathayana Mn treatment resulted in oxidative stress indicated by the oxidation to lipids, proteinsand DNA. A regulated network of defence strategies was employed for the chelation,detoxification and tolerance of Mn including the enhanced synthesis of ABA and polyamines,the accumulation of free amino acids, especially His and Pro, and the activation of theenzymes superoxide dismutase and guaiacol peroxidase. Contents of non-protein thiol,reduced glutathione, phytochelatins and phenolics compounds and activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase also increased significantly forantioxidant or chelation functions. The wet climate population not only accumulated lessabscisic acid, free amino acids, phytochelatins and phenolics compounds, but also exhibitedlower activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase thusresulting in more serious oxidative damage and more curtained growth.
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克隆植物被认为比非克隆植物更宜于利用异质性环境。在复杂的空间异质环境中,克隆植物可能形成了各种有效利用环境异质性的适应对策。对于克隆植物适应机制的研究,前人已做了大量的工作,特别是从形态和生物量分配等方面对简单异质生境下克隆植物的克隆整合和克隆分工进行了详细的研究。本研究以分布广泛的克隆植物东方草莓(Fragaria orientalis)作为研究对象,应用野外调查和实验生态学方法,采用多对比度单资源模型和不同向双资源模型,从形态和生理生态的角度,研究复杂异质生境下克隆植物的整合和分工及其耗益问题,分析不同类型的生境对克隆植物整合和分工的修饰作用,进而探讨克隆植物对异质生境的适应策略。克隆构型和分株种群特征是植物克隆生长及其生态适应对策研究的基本内容。本文通过野外调查,研究在不同光照条件下东方草莓克隆构型、分株种群特征以及点分布格局。结果表明:东方草莓的克隆构型随光照发生相应的变化,低光照下其匍匐茎节间长和分枝角度均增大而分枝强度减小;随光照减弱,东方草莓分株种群的生物量、根冠比和分株种群密度显著降低;不同光照下东方草莓分株均以随机分布为主但不同尺度下有所差异,其分布格局强度依次为旷地<林缘<林下。结合克隆植物对资源的利用对策,探讨了克隆构型和分株种群特征以及分布格局随环境条件变化的生态适应意义。不同生境斑块条件下克隆植物可能采取不同的适应对策。采用盆栽实验,研究不同水分对比度下克隆整合及其生理生态特征,并对单向和交互资源中东方草莓的克隆整合做了对比研究。结果显示:高的水分对比度能够促进东方草莓的克隆整合,并能刺激相连分株增加光合作用,东方草莓体内的氧化—抗氧化系统也II随对比度做出相应的反应。耗-益分析表明胁迫分株的受益是以供给分株的损耗为代价的,但从克隆片段总体来说是受益的。单向资源中东方草莓生长的绝对值高于交互资源,但耗-益分析表明生长于交互资源下东方草莓的克隆整合获益大于生长于单向资源下东方草莓的克隆整合获益。长期生长于特定生境的克隆植物,在进化过程中其克隆整合和克隆分工在对资源异质性的适应策略方面可能有所侧重。采用盆栽实验对来自不同海拔梯度的东方草莓的克隆整合和克隆分工对异质资源的适应对策进行了研究。实验结果表明,来自高海拔的东方草莓可塑性较差。来自两个海拔的东方草莓对切断匍匐茎的表现有所差异,总体上切断匍匐茎对来自高海拔的东方草莓影响更大些。另外,来自高海拔的东方草莓表现出更高的克隆分工。IIIClonal plants are known to be more suitable for the habitats of heterogeneousresources than nonclonal plants, perhaps due to their well developed adaptivestrategies to environmental heterogeneity. Many studies have been done on theadaptive mechanisms of clonal plants, especially on the clonal integration anddivision of labor with morphology and biomass allocation under simpleheterogeneous habitats. Based on field surveys, laboratory experiments, multi-contrastunidirectional resource model and reciprocal resource model, Fragaria orientalis, aRosaceae stoloniferous herb that widely distributes in China, was used to study thisplant’s morphological and physiological responses to complicated heterogeneoushabitats in terms of its clonal integration, division of labor and cost-benefit, as well astheir modifications by different habitats, so as to better understand the adaptivestrategies of clonal plants under heterogeneous environments.Clonal architecture and ramet population characteristics are of the major concernin the studies on growth and adaptive strategies of clonal plants. Clonal architecture,ramet population characteristics and spatial point pattern of F. orientalis underdifferent light intensity were studied with field observations. The results showed that,clonal architecture changed with light availability: Internode-lengths and branchangels of stolons were larger while branch intensities were smaller under lower lightintensity than those under higher light intensity; Biomass of ramet population,root-shoot ratio and density of ramet population decreased significantly with reduce oflight intensity; Under all light intensities, spatial pattern of ramets was mainlyrandomly distributed but it changed with different scales, with pattern intensity as:open space < forest edge < understory. Adaptation significance of the clonal architecture, the ramet population characteristics and the spatial pattern changing withdifferent environments was discussed according to these results.Clonal plants may take different adaptive strategies under different patches. Withpot culture, clonal integration and physiological parameters of F. orientalis underdifferent water contrasts were studied, and clonal integration under unilateralresources and reciprocal resources were also compared. The results suggested that,high water contrast improve the clonal integration of F. orientalis and increase thephotosynthesis of connected ramets. Oxidative and antioxidative system of F.orientalis also responded with changing water contrasts. According to cost-benefitanalysis, the drought-stressed ramets obtained benefits from the connectedwell-watered ramets, and as a whole, the clonal fragment could also get benefits.Growth of F. orientalis in homogeneous resources was better than that inheterogeneous resources, but the whole plant got more benefit through clonalintegration in heterogeneous resources than in homogeneous resources.Pot culture experiments were also used to study the adaptive strategies inutilizing heterogeneous resources by the plant populations from different altitudes.The results showed that, F. orientalis from alpine zones were shorter and lessexpanded with poorer clonal plasticity than those from middle mountains. F.orientalis from two different altitudes showed different responses to stolon severing,and as a whole, stolon severing had more influence on F. orientalis from alpine zones.In addition, F. orientalis from alpine zones exhibited higher division of labor, whichsuggested that clonal plants from different habitats develop their own adaptivemechanisms in their clonal integration and division of labor in response toenvironmental heterogeneity.
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小鲵属为亚洲特有的有尾两栖类,是小鲵科之模式属。现记载小鲵属动物有29种,占全科物种数一半以上(Frost, 2007),为小鲵科第一大属。该属分布跨越古北界和东洋界,分布于中国、朝鲜、韩国、日本等地区,其系统学研究一直以来颇为中外学者所关注。澄清该属的物种分类问题,阐明其种间的系统发育关系对整个小鲵科的系统演化与分布格局关系的研究具有关键性意义。 本论文以中国及周边地区的小鲵属物种为主要对象,主要利用分子生物学实验与生物信息学途径相结合的手段,运用支序系统学与分子进化生物学理论及分析方法,展开系统发育的研究。在此基础上诠释现存的分类问题,并探讨该属系统发育关系。 研究材料上,本研究采用野外采集与网络下载数据相结合的方法,获取了较为全面的小鲵属物种DNA序列资料。技术手段上,选取了线粒体DNA的Cytb、12S、16S、NADH 2、COI等多个基因部分片段序列,对小鲵属开展了较为全面系统的研究。分析方法上,针对小鲵属物种各类群的具体情况,运用了处于领域前沿的多种分析方法。应用PAUP、MrBayes、Modeltest、Mega等软件,采用了最大简约法(MP)、邻接法(NJ)、贝叶斯推断(BI)及K2P遗传距离分析等方法。 本研究对小鲵属进行了较为全面的系统发育研究,弥补了有关小鲵属系统发育研究的不足,并得出了以下结果: (1)关于豫南小鲵Hynobius yunanicus的有效性,基于细胞色素b序列的系统发育关系联合形态和染色体组型等证据证明了豫南小鲵是商城肥鲵的同物异名。 (2)获得了较为全面的小鲵属物种系统发育树,并以此解释了北海道滞育小鲵、东北小鲵、中国小鲵与义乌小鲵等存在的分类问题。 (3)本研究利用DNA条形码技术对小鲵属及小鲵科物种进行了鉴定,再次证明豫南小鲵为商城肥鲵的同物异名;并认为猫儿山小鲵与挂榜山小鲵为同物异名。 综上,本研究较为完整地勾勒了小鲵属的系统发育关系全貌,并对小鲵属物种的起源进行了推测。 Hynobius, the type genus of the Family Hynobiidae, is the only exclusively Asian salamander genus. This genus which contains 29 species (beyond half of total Family), is the key group in Hynobiidae. The genus distributed across Palaearctic and Oriental Realm, and was found in China, Korea, and Japan. Systematics of genus Hynobius draws attention of researchers all the times. Resolving the taxonomic and phynogenetic questions of Hynobius is very important to the evolutionary research of Family Hynobiidae. Firstly, studies on systematics of genus Hynobius based on morphology, karyotype and molecular phylogeny of Hynobius are reviewed along with existing questions of this genus. The sequential reaserch project of phylogenetics is perspectively outlined. Using molecular data, we compared Hynobius yunanicus with a sympatric species Pachyhynobius shangchengensis. Our cytb sequences associating with karyotypic and morphological data supportted that H. yunanicus is not a valid species, but a synonym of P. shangchengensis. Because of phenotypic plasticity, some morphological characters are not even suitable for identifying hynobiids. The taxonomy of hynobiids is still controversial to a certain extent (Zhao et al. 1993; Fei, 1999; Chen et al. 2001; Zeng et al. 2006) and needs to be resolved by a new method. Here we examined the utility of COI barcoding for the discrimination of hynobiids. Meantime, the taxonomy of this Family was looked-over again. Our result show that the DNA Barcoding based on COI is easier and more rapidly than classic methods. And the DNA Barcodes data supported the actual taxonomy of Hynobiidae. Based on the achievements of our research, the phylogeny of Hynobius was reconstructed including some new species (H. maoershanensis, H. guabangshanensis, etc). Besides the phylogenetics of Hynobius was outlined, some questions and the hypothesis about the origin of genus Hynobius was put out.
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p21(Waf1/Cip1), best known as a broad-specificity inhibitor of cyclin/cyclin-dependent kinase complexes, can interact with various target proteins, and this ability relies on its structural plasticity. Therefore, studies on the structural properties of p(21) are very important to understand its structure-function relationship. However, detailed studies on its secondary structure and biophysical properties have been comparatively sparse. A human p(21) gene was cloned into the temperature expression vector pBV220 and transformed into Escherichia coli strain JM109.
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Circadian growth rhythm of the juvenile sporophyte of the brown alga Undaria pirznatifida was measured with the computer-aided image analysis system in constant florescent white light under constant temperature (10 degrees C). The growth rhythm persisted for 4 d in constant light with a free-running period of 25. 6 h. Egg release from filamentous gametophytes pre-cultured in the light - dark regime was evaluated for six consecutive days at fixed time intervals in constant white light and 12 h light per day. Egg release rhythm persisted for 3 d in both regimes, indicating the endogenous nature. Temporal scale of egg release and gametogenesis in 18, 16, 12 and 8 h light per day were evaluated respectively using vegetatively propagated filamentous gametophytes. Egg release occurred 2 h after the onset of dark phase and peaked at midnight. Evaluation of the rates of oogonium formation, egg release or fertilization revealed no significant differences in four light-dark regimes, indicating; the great plasticity of sexual reproduction. No photoperiodic effect in gametogenesis in terms of oogonium formation and egg release was found, but fertilization in short days was significantly higher than in long days. Results of this investigation further confirmed the general occurrence of circadian rhythms in inter-tidal seaweed species.
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Photoinhibition is a central problem for the understanding of plasticity in photosynthesis vs. irradiance response. It effectively reduces the photosynthetic rate. In this contribution, we present a mechanistic model of algal photoinhibition induced by photodamage to photosystem-II. Photosystem-IIs (PSIIs) are assumed to exist in three states: open, closed and inhibited. Photosynthesis is closely associated with the transitions between the three states. The present model is defined by four parameters: effective cross section of PSII, number of PSIIs, turnover time of electron transfer chains and the ratio of rate constant of damage to that of repair of D1 proteins in PSIIs. It gives a photosynthetic response curve of phytoplankton to irradiance (PI-curve). Without photoinhibition, the PI-curve is in hyperbola with the first three parameters. The PI-curve with photoinhibition can be simplified to the same form as the hyperbola by replacing either the number of PSIIs with the number of functional PSIIs or the turnover time of electron transfer chains with the average turnover time.
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Expansive soil is a kind of typical unsaturated soil with characteristics of high swelling-shrinking deformation, cracks and over consolidation. It is very harmful to civil engineering, As a new processing method deal with expansive soil, Chemistry treatment has widespread applied in developed countries such as Europe and America, and also gained remarkable result. Based on the embankment filling soil improving testing projects in Meng-Xin freeway, this paper proposed a new processing method to expansive soil embankment wrapped with PAS-treated soil, experimental study of expansive soil chemical improved by PAS is been carried out. The water content change is the external factor which causes expansive soil to have swelling-shrinkage deformation. this reflected that the soil body swelling-shrinkage characteristic mainly depends on its mineral ingredient and the soil-water mutual function. This paper takes expansive soil as one kind of ordinary high plastic clay from angle of clay-water mutual function explained the expansive soil swelling-shrinkage deformation mechanism on microscopic. And take this swelling-shrinkage mechanism as the master line, Cooperates with the China Academy of Chemistry, we developed the new method PAS treatment, trough ionic exchange, joint, package and flocculation, the stronger static electricity function weakened the level through adsorption and the stronger static electricity function, PAS can weakened the negative charge repulsion between levels, causes the electric potential to reduce, diffusion layer thickness to be thinner, and improves the water affinity performance of expansive soil effectively. Moreover the space network architecture compromised with PAS and soil enhanced the joint strength between the clay particles , enable the soil body to have comparatively high strength and the distortion rate. pointed proposed the PAS modified principle. Combine with the construction of experimented road, this paper sums up and presents the construction craft and technology requirement of PAS treatment to expansive soil embankment. Through many experimental studied the basic physical property, the intensity characteristic and water stability changes of expansive soil and PAS-treated soil. The results of study indicate that adding lime into the expansive soil can reduce the content of clay gain obviously, reduce the plasticity notably, increase the strength greatly, control the property of swelling and shrinking effectively, and can meliorate the stability of sucking water clearly. Simultaneity PAS don’t change the cultivate capacity of the soil, the modified slope of the embankment can adopt plant fixed slope method as ecology protection. Finally the processing effect of use different treatment has analyzed through numerical simulation, summarized the PAS chemical wrapping treatment process in the actual project application, and appraised its processing effect and the project efficiency. The research indicated that PAS chemical treatment is one effective method to improve expansive soil. Compare with long-distance replacement, especially in the high plastic expansive soil massive distribution area, PAS treatment has the very greatly economical superiority to be promoted. The study in the paper not only afforded technique method to Meng-Xin expressway construction but also important for improvement of the expressway construction theory in swelling soil areas. Key words: PAS; expansive soil; swelling-shrinkage deformation mechanism; wrapping embankment; chemical modified treatment.
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Based on the features of soft soil in Tianjing Coastal New Developing Area, this kind of soil with different content of sand was researched systematically, according to the indoor experiment, about its characteristics of strength and deformation. The main results are summarized in the following: Firstly, on the basis of geological engineering investigation, the systemic experiments about the physical characteristics were conducted. The test soil samples were taken from the gray and gray-yellow silty soft soil which was formatted by near-shore marine sediment and marine-continental interactive sediment. The original condition of the sample soil was in saturation and the basic indexes are: liquid limit36.1%, plastic limit 18.8%, plasticity index. Then, the condensation characteristics of the soft soil were analyzed through high-pressure consolidation tests. The results show that,in various loading serials, the coefficient of compressibility under P=100kPa and 200kPa are all larger than 0.5MPa-1. So the sample soil is a kind of high-compressibility soil. Secondly, triaxial strength of undisturbed soil and remoulded soil was researched by using triaxial test. The types of stress-strain curve of both undisturbed and remoulded soil are the stress stabilization and softening type, which show the specific plastic character. Furthermore, the cohesion and friction angle of undisturbed soil changes, when the ambient pressure is different, instead of a stable value for all time; the cohesion and friction angle of remoulded soil changes with the compactness and sand-carrying capacity which is wholly higher than undisturbed soil. At last, the stress-strain results of both undisturbed and remoulded soil were normalized by using the ambient pressure as the normalization factor. The results show that, there are all some normalization characters in both undisturbed and remoulded soil, however, the feature of normalization of undisturbed soil is worse than the remoulded ones. The main reason is that the undisturbed samples are worse in equality and the unavoidable disturb through the process of sampling and experiments will also make them can not put up good normalization. Therefore, it is feasible to normalize the soil in Tianjing Coastal New Developing Area with the ambient pressure as normalization factor.
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The unsaturated expansive soil is a hotspot and difficulty in soil mechanics inland and outland. The expansive soil in our China is one of the widest in distributing and greatest in area, and the disaster of expansive soil happens continually as a result. The soil mechanics test, monitor, numerical simulation and engineering practice are used to research swell and shrinkage characteristic, edge strength characteristic and unsaturated strength characteristic of Mengzi expansive soil. The seep and stability of the slope for expansive soil associated with fissure are analyzed and two kinds of new technique are put forward to be used in expansive soil area, based on disaster mechnics proposed of the slope.The technique of reinforcement in road embankment is optimized also. Associated with engineering geology research of Mengzi expansive soil, mineral composition, chemical composition, specific area and cation content, dissolubility salt and agglutinate, microcosmic fabric characteristic, cause of formation and atmosphere effect depth are analyzed to explain the intrinsic cause and essence of swell and shrinkage for expansive soil. The rule between swell-shrinkage and initial state, namely initial water content, initial dry density and initial pressure, can be used to construction control. Does Response model is fit to simulate the rule, based on ternary regression analysis. It has great meaning to expansive soil engineering in area with salt or alkali. The mechanics under CD, CU and GCU of expansive soil is researched by edge surface theory to explain the remarkable effect of consolidation pressure, initial dry density, initial water content, cut velocity, drainage and reinforcement to the edge strength characteristic. The infirm hardening stress strain curves can be fitted with hyperbola model and the infirm softening curves can be fitted with exponential model. The normalization theory can be used to reveal the intrinsic unity of the otherness which is brought by different methods to the shear strength of the same kinds of samples. The unsaturated strain softening characteristic and strength envelope of remolding samples are researched by triaxial shear test based on suction controlled, the result of which is simulated by exponential function. The strength parameters of the unsaturated samples are obtained to be used in the unsaturated seep associated with rainfall. The elasticity and plasticity characters of expansive soil are researched to attain the model parameters by using modified G-A model. The humidification destroy characteristic of expansive soil is discussed to research the disaster mechanism of the slope with the back pressure increasing and suction decreasing under bias pressure consolidation. The indoor and outdoor SWCCs are measured to research the effect factors and the rule between different stress and filling environment. The moisture absorption curves can express the relationship between suction and water content in locale. The SWCCs of Mengzi expansive soil are measured by GDS stress path trixial system. The unsaturated infiltration function is gained to research seep and stability of the slope of expansive soil. The rainfall infiltration and ability of slope considering multifarious factors are studied by analyzing fissure cause of Mengzi expansive soil. The mechanism of the slope disaster is brought forward by the double controlling effect between suction and fissure. Two new kinds of technique are put forward to resolve disaster of expansive soil and the technique of reinforcement on embankment is optimized, which gives a useful help to solving engineering trouble.
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Rock mass is widely recognized as a kind of geologic body which consists of rock blocks and discontinuities. The deformation and failure of rock mass is not only determined by rock block,but also by discontinuity which is virtually more important. Mutual cutting and combination of discontinuities controlled mechanical property of rock mass. The complex cutting of discontinuities determine the intense anisotropy on mechanical property of rock mass,especially under the effect of ground stress. Engineering practice has show that the brittle failure of hard rock always occurs when its working stress is far lower than the yield strength and compressive strength,the failure always directly related to the fracture propagation of discontinuities. Fracture propagation of discontinuities is the virtue of hard rock’s failure. We can research the rock mass discontinuous mechanical properties precisely by the methods of statistical analysis of discontinuities and Fracture Mechanics. According to Superposition Principle in Fracture Mechanics,A Problem or C Problem could be chosen to research. Problem A mainly calculates the crack-tip stress field and displacement field on internal discontinuities by numerical method. Problem C calculate the crack-tip stress field and displacement field under the assumption of that the mainly rock mass stress field has been known. So the Problem C avoid the complex mutual interference of stress fields of discontinuities,which is called crack system problem in Fracture Mechanics. To solve Problem C, field test on stress field in the rock mass is needed. The linear Superposition of discontinuities strain energies are Scientific and Rational. The difference of Fracture Mechanics between rock mass and other materials can mostly expression as:other materials Fracture Mechanics mostly face the problem A,and can’t avoid multi-crack puzzle, while the Rock mass Fracture Mechanics answer to the Problem C. Problem C can avoid multi-discontinuities mutual interference puzzle via the ground stress test. On the basis of Problem C, Fracture Mechanics could be used conveniently in rock mass. The rock mass statistics fracture constitutive relations, which introduced in this article, are based on the Problem C and the Discontinuity Strain Energy linear superposition. This constitutive relation has several merits: first, it is physical constitutive relation rather than empirical; second, it is very fit to describe the rock mass anisotropy properties; third, it elaborates the exogenous factors such as ground stress. The rock mass statistics fracture constitutive relation is the available approach to answer to the physical, anisotropic and ground stress impacted rock mass problems. This article stand on the foundation of predecessor’s statistics fractures constitutive relation, and improved the discontinuity distributive function. This article had derived the limitation of negative exponential distribution in the course of regression analysis, and advocated to using the two parameter negative exponential distribution for instead. In order to solve the problems of two-dimension stability on engineering key cross-sectional view in rock mass, this article derived the rock mass planar flexibility tensor, and established rock mass two-dimension penetrate statistics fracture constitutive relation on the basis of penetrate fracture mechanics. Based on the crack tip plasticity research production of penetrate fracture, for example the Irwin plasticity equifinality crack, this article established the way to deal with the discontinuity stress singularity and plastic yielding problem at discontinuity tip. The research on deformation parameters is always the high light region of rock mass mechanics field. After the dam foundation excavation of XiaoWan hydroelectric power station, dam foundation rock mass upgrowthed a great deal of unload cracks, rock mass mechanical property gotten intricacy and strong anisotropy. The dam foundation rock mass mostly upgrowthed three group discontinuities: the decantation discontinuity, the steep pitch discontinuity, and the schistosity plane. Most of the discontinuities have got partial unload looseness. In accordance with ground stress field data, the dam foundation stress field greatly non-uniform, which felled under the great impaction of tectonic stress field, self-weight stress field, excavation geometric boundary condition, and excavation, unload. The discontinuity complexity and stress field heterogeneity, created the rock mass mechanical property of dam foundation intricacy and levity. The research on the rock mass mechanics, if not take every respected influencing factor into consideration as best as we can, major errors likely to be created. This article calculated the rock mass elastic modulus that after Xiao Wan hydroelectric power station dam foundation gutter excavation finished. The calculation region covered possession monolith of Xiao Wan concrete double-curvature arch dam. Different monolith were adopted the penetrate fracture statistics constitutive relation or bury fracture statistics constitutive relation selectively. Statistics fracture constitutive relation is fit for the intensity anisotropy and heterogeneity rock mass of Xiao Wan hydroelectric power station dam foundation. This article had contrastive analysis the statistics fracture constitutive relation result with the inclined plane load test actual measurement elastic modulus and RMR method estimated elastic modulus, and find that the three methods elastic modulus have got greatly comparability. So, the statistics fracture constitutive relations are qualified for trust. Generally speaking,this article had finished following works based on predecessors job: “Argumentation the C Problems of superposition principle in Fracture Mechanics, establish two-dimension penetrate statistics fracture constitutive relation of rock mass, argue the negative exponential distribution limitation and improve it, improve of the three-dimension berry statistics fracture constitutive relation of rock mass, discontinuity-tip plastic zone isoeffect calculation, calculate the rock mass elastic modulus on two-dimension cross-sectional view”. The whole research clue of this article inherited from the “statistics rock mass mechanics” of Wu Faquan(1992).
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The research on mechanical effects of water-rock and soil interaction on deformation and failure of rocks and soils involves three aspects of mechanics, physics and chemistry. It is the cross between geochemistry and rock mechanics and soil mechanics. To sum up, the mechanical effects of water-rock and soil interaction is related to many complex processes. Research in this respect has been being an important forward field and has broad prospects. In connection with the mechanism of the effects of the chemical action of water-rock on deformation and failure of rocks and soils, the research significance, the present state, the developments in this research domain are summarized. Author prospects the future of this research. The research of the subject should be possessed of important position in studying engineering geology and will lead directly to a new understand on geological hazard and control research. In order to investigation the macroscopic mechanics effects of chemical kinetics of water-rock interaction on the deformation and failure, calcic rock, red sandstone and grey granite reacting chemically with different aqueous solution at atmospheric temperature and atmospheric pressure are uniaxially compressed. The quantitative results concerning the changes of uniaxially compressive strength and elastic modulus under different conditions are obtained. It is found that the mechanical effects of water on rock is closely related to the chemical action of water-rock or the chemical damage in rock, and the intensity of chemical damage is direct ratio to the intensity of chemical action in water-rock system. It is also found that the hydrochemical action on rock is time-dependent through the test. The mechanism of permeation and hydrochemical action resulting in failure of loaded rock mass or propagation of fractures in rocks would be a key question in rock fracture mechanics. In this paper, the fracture mechanical effects of chemical action of water-rock and their time- and chemical environment-dependent behavior in grey granite, green granite, grey sandstone and red sandstone are analyzed by testing K_(IC) and COD of rock under different conditions. It is found that: ①the fracture mechanical effect of chemical action of water-rock is outstanding and time-dependent, and high differences exist in the influence of different aqueous solution, different rocks, different immersion ways and different velocity of cycle flow on the fracture mechanical effects in rock. ②the mechanical effects of water-rock interaction on propagation of fractures is consistent with the mechanical effects on the peak strength of rock. ③the intensity of the mechanical fracture effects increases as the intensity of chemical action of water-rock increases. ④iron and calcium ion bearing mineral or cement in rock are some key ion or chemical composition, and especially iron ion-bearing mineral resulting in chemical action of water-rock to be provided with both positive and negative mechanical effects on rock. Through the above two tests, we suggest that primary factors influencing chemical damage in rock consist of the chemical property of rock and aqueous solution, the structure or homogeneity of rocks, the flow velocity of aqueous solution passing through rock, and cause of formation or evolution of rock. The paper explores the mechanism on the mechanical effects of water-rock interaction on rock by using the theory of chemistry and rock fracture mechanics with chemical damage proposed by author, the modeling method and the energy point of view. In this paper, the concept of absorbed suction between soil grains caused by capillary response is given and expounded, and the relation and basic distinction among this absorbed suction, surface tension and capillary pressure of the soil are analyzed and established. The law of absorbed suction change and the primary factors affecting it are approached. We hold that the structure suction are changeable along with the change of the saturation state in unsaturated soils. In view of this, the concept of intrinsic structure suction and variable structure suction are given and expounded, and this paper points out: What we should study is variable structure suction when studying the effective stress. By IIIy κHH's theory of structure strength of soils, the computer method for variable structure suction is analyzed, the measure method for variable structure suction is discussed, and it reach the conclusions: ①Besides saturation state, variable structure suction is affected by grain composition and packing patter of grains. ②The internal relations are present between structure parameter N in computing structure suction and structure parameter D in computing absorbed suction. We think that some problems exit in available principle of effective stress and shear strength theory for unsaturated soil. Based on the variable structure suction and absorbed suction, the classification of saturation in soil and a principle of narrow sense effective stress are proposed for unsaturated soils. Based on generalized suction, the generalized effective stress formula and a principle of generalized effective stress are proposed for unsaturated soils. The experience parameter χ in Bishop's effective stress formula is defined, and the principal factors influencing effective stress or χ. The primary factor affecting the effective stress in unsaturated soils, and the principle classifying unsaturated soils and its mechanics methods analyzing unsaturated soils are discussed, and this paper points out: The theory on studying unsaturated soil mechanics should adopt the micromechanics method, then raise it to macromechanics and to applying. Researching the mechanical effects of chemical action of water-soil on soil is of great importance to geoenvironmental hazard control. The texture of soil and the fabric of soil mass are set forth. The tests on physical and mechanical property are performed to investigate the mechanism of the positive and negative mechanical effects of different chemical property of aqueous solution. The test results make clear that the plastic limit, liquid limit and plasticity index are changed, and there exists both positive and negative effects on specimens in this test. Based on analyzing the mechanism of the mechanical effects of water-soil interaction on soil, author thinks that hydrochemical actions being provided with mechanical effects on soil comprise three kinds of dissolution, sedimentation or crystallization. The significance of these tests lie in which it is recognized for us that we may improve, adjust and control the quality of soils, and may achieve the goal geological hazard control and prevention.The present and the significance of the research on environmental effects of water-rock and soil interaction. Various living example on geoenvironmental hazard in this field are enumerated. Following above thinking, we have approached such ideals that: ①changing the intensity and distribution of source and sink in groundwater flow system can be used to control the water-rock and soil interaction. ②the chemical action of water-rock and soil can be used to ameliorate the physical and mechanical property of rocks and soils. Lastly, the research thinking and the research methods on mechanical effects and environmental effects of water-rock and soil interaction are put forward and detailed.